How does moss photosynthesis relate to leaf and canopy structure? Trait relationships for 10 Hawaiian species of contrasting light habitats.

Mosses are an understudied group of plants that can potentially confirm or expand principles of plant function described for tracheophytes, from which they diverge strongly in structure. We quantified 35 physiological and morphological traits from cell-, leaf- and canopy-level, for 10 ground-, trunk- and branch-dwelling Hawaiian species. We hypothesized that trait values would reflect the distinctive growth form and slow growth of mosses, but also that trait correlations would be analogous to those of tracheophytes. The moss species had low leaf mass per area and low gas exchange rate. Unlike for tracheophytes, light-saturated photosynthetic rate per mass (A(mass)) did not correlate with habitat irradiance. Other photosynthetic parameters and structural traits were aligned with microhabitat irradiance, driving an inter-correlation of traits including leaf area, cell size, cell wall thickness, and canopy density. In addition, we found a coordination of traits linked with structural allocation, including costa size, canopy height and A(mass). Across species, A(mass) and nitrogen concentration correlated negatively with canopy mass per area, analogous to linkages found for the 'leaf economic spectrum', with canopy mass per area replacing leaf mass per area. Despite divergence of mosses and tracheophytes in leaf size and function, analogous trait coordination has arisen during ecological differentiation.

[1]  Jill,et al.  Desiccation Tolerance , 2012 .

[2]  L. Sack,et al.  Leaf Trait Diversification and Design in Seven Rare Taxa of the Hawaiian Plantago Radiation , 2009, International Journal of Plant Sciences.

[3]  D. Hanson,et al.  Do bryophyte shoot systems function like vascular plant leaves or canopies? Functional trait relationships in Sphagnum mosses (Sphagnaceae). , 2008, American journal of botany.

[4]  T. Giambelluca,et al.  Scaling of Frond Form in Hawaiian Tree Fern Cibotium glaucum: Compliance with Global Trends and Application for Field Estimation , 2008 .

[5]  Huayun Xiao,et al.  Tissue N content and 15N natural abundance in epilithic mosses for indicating atmospheric N deposition in the Guiyang area, SW China , 2008 .

[6]  H. Griffiths,et al.  To concentrate or ventilate? Carbon acquisition, isotope discrimination and physiological ecology of early land plant life forms , 2008, Philosophical Transactions of the Royal Society B: Biological Sciences.

[7]  Thomas J. Givnish,et al.  COMPARATIVE STUDIES OF LEAF FORM: ASSESSING THE RELATIVE ROLES OF SELECTIVE PRESSURES AND PHYLOGENETIC CONSTRAINTS , 2008 .

[8]  Lars Markesteijn,et al.  Seedling Traits Determine Drought Tolerance of Tropical Tree Species , 2008 .

[9]  R. Mäkipää,et al.  Differences in the growth response of three bryophyte species to nitrogen. , 2008, Environmental pollution.

[10]  J. Bennetzen,et al.  The Physcomitrella Genome Reveals Evolutionary Insights into the Conquest of Land by Plants , 2008, Science.

[11]  David Johnson,et al.  Long-term nitrogen deposition increases phosphorus limitation of bryophytes in an ombrotrophic bog , 2008, Plant Ecology.

[12]  D. Wardle,et al.  Ecosystem input of nitrogen through biological fixation in feather mosses during ecosystem retrogression , 2007 .

[13]  G. Zotz,et al.  A moss “canopy” – Small-scale differences in microclimate and physiological traits in Tortula ruralis , 2007 .

[14]  R. Ozolinčius,et al.  Wood ash and nitrogen influence on ground vegetation cover and chemical composition , 2007 .

[15]  A. Wood,et al.  Desiccation-tolerance in bryophytes: a review , 2007 .

[16]  Peter B Reich,et al.  The scaling of leaf area and mass: the cost of light interception increases with leaf size , 2007, Proceedings of the Royal Society B: Biological Sciences.

[17]  F. Valladares,et al.  Do we underestimate the importance of leaf size in plant economics? Disproportional scaling of support costs within the spectrum of leaf physiognomy. , 2007, Annals of botany.

[18]  A. Britton,et al.  NP stoichiometry of low-alpine heathland: Usefulness for bio-monitoring and prediction of pollution impacts , 2007 .

[19]  F. Valladares,et al.  2 Opportunistic Growth and Desiccation Tolerance: The Ecological Success of Poikilohydrous Autotrophs , 2007 .

[20]  T. Brodribb,et al.  Leaf Maximum Photosynthetic Rate and Venation Are Linked by Hydraulics1[W][OA] , 2007, Plant Physiology.

[21]  B. Mishler,et al.  Bryophyte phylogeny: Advancing the molecular and morphological frontiers , 2007 .

[22]  Ian J. Wright,et al.  “Diminishing returns” in the scaling of functional leaf traits across and within species groups , 2007, Proceedings of the National Academy of Sciences.

[23]  Nadejda A. Soudzilovskaia,et al.  Comparative cryptogam ecology: a review of bryophyte and lichen traits that drive biogeochemistry. , 2007, Annals of botany.

[24]  Nadejda A. Soudzilovskaia,et al.  Comparative Ecology and Ecological Scaling Highlight: Article , 2007 .

[25]  M. Lechowicz,et al.  Are correlations among foliar traits in ferns consistent with those in the seed plants? , 2007, The New phytologist.

[26]  S. Gower,et al.  Estimation of stand-level leaf area for boreal bryophytes , 2007, Oecologia.

[27]  J. Frahm,et al.  Historical alteration in the nitrogen concentration and 15N natural abundance of mosses in Germany: Indication for regionally varying changes in atmospheric nitrogen deposition within the last 140 years , 2006 .

[28]  V. Lieffers,et al.  Dynamics of mineral nitrogen released from feathermosses after dehydration or handling stress , 2006 .

[29]  R. Quatrano,et al.  Mosses as model systems for the study of metabolism and development. , 2006, Annual review of plant biology.

[30]  M. Westoby,et al.  Bivariate line‐fitting methods for allometry , 2006, Biological reviews of the Cambridge Philosophical Society.

[31]  P. Reich,et al.  Fundamental trade-offs generating the worldwide leaf economics spectrum. , 2006, Ecology.

[32]  Ü. Niinemets,et al.  Structural determinants of leaf light-harvesting capacity and photosynthetic potentials , 2006 .

[33]  Lawren Sack,et al.  Leaf hydraulics. , 2006, Annual review of plant biology.

[34]  F. Putz,et al.  Ecophysiology in Relation to Exposure of Pendant Epiphytic Bryophytes in the Canopy of a Tropical Montane Oak Forest 1 , 2005 .

[35]  B. Mishler,et al.  Desiccation Tolerance in Bryophytes: A Reflection of the Primitive Strategy for Plant Survival in Dehydrating Habitats?1 , 2005, Integrative and comparative biology.

[36]  M. Proctor Why do Polytrichaceae have lamellae? , 2005 .

[37]  T. Green,et al.  Photosynthetic responses of three common mosses from continental Antarctica , 2005, Antarctic Science.

[38]  C. W. Smith,et al.  A revised checklist of Hawaiian mosses , 2004 .

[39]  K. Hikosaka Interspecific difference in the photosynthesis–nitrogen relationship: patterns, physiological causes, and ecological importance , 2004, Journal of Plant Research.

[40]  M. Proctor,et al.  Are bryophytes shade plants? Photosynthetic light responses and proportions of chlorophyll a, chlorophyll b and total carotenoids. , 2004, Annals of botany.

[41]  A. Dijk,et al.  The importance of epiphytes to total rainfall interception by a tropical montane rain forest in Costa Rica , 2004 .

[42]  J. Osborne,et al.  Sample size and subject to item ratio in principal components analysis. , 2004 .

[43]  Sean C. Thomas,et al.  The worldwide leaf economics spectrum , 2004, Nature.

[44]  Rebecca A Montgomery,et al.  Adaptive radiation of photosynthetic physiology in the Hawaiian lobeliads: light regimes, static light responses, and whole-plant compensation points. , 2004, American journal of botany.

[45]  David D. Ackerly,et al.  FUNCTIONAL STRATEGIES OF CHAPARRAL SHRUBS IN RELATION TO SEASONAL WATER DEFICIT AND DISTURBANCE , 2004 .

[46]  W. Oechel,et al.  Moss functioning in different taiga ecosystems in interior Alaska , 1981, Oecologia.

[47]  I. Ohad,et al.  Physcomitrella patens and Ceratodon purpureus, mosses as model organisms in photosynthesis studies , 2004, Photosynthesis Research.

[48]  N. Holbrook,et al.  The ‘hydrology’ of leaves: co‐ordination of structure and function in temperate woody species , 2003 .

[49]  A. J. Shaw Structural Diversity of Bryophytes , 2003 .

[50]  R. Gabriel,et al.  Responses of photosynthesis to irradiance in bryophytes of the Azores laurel forest , 2003 .

[51]  J. Hicke,et al.  Global synthesis of leaf area index observations: implications for ecological and remote sensing studies , 2003 .

[52]  P. Reich,et al.  The Evolution of Plant Functional Variation: Traits, Spectra, and Strategies , 2003, International Journal of Plant Sciences.

[53]  M. A T T H E,et al.  The contribution of bryophytes to the carbon exchange for a temperate rainforest , 2003 .

[54]  Z. Tuba,et al.  Poikilohydry and homoihydry: antithesis or spectrum of possibilities? , 2002, The New phytologist.

[55]  G. Zotz,et al.  Seasonal Changes in Diel CO2 Exchange of Three Central European Moss Species: a One-Year Field Study , 2001 .

[56]  H. Crum Structural Diversity of Bryophytes , 2001 .

[57]  Ülo Niinemets,et al.  GLOBAL-SCALE CLIMATIC CONTROLS OF LEAF DRY MASS PER AREA, DENSITY, AND THICKNESS IN TREES AND SHRUBS , 2001 .

[58]  C. Martin,et al.  Photosynthetic capacity of mosses relative to vascular plants , 2001 .

[59]  M. Proctor,et al.  Mosses and alternative adaptation to life on land. , 2000, The New phytologist.

[60]  D. Cove The Moss, Physcomitrella patens , 2000, Journal of Plant Growth Regulation.

[61]  D. Vitt Bryophyte Biology: Peatlands: ecosystems dominated by bryophytes , 2000 .

[62]  Ying-di Liu,et al.  The measurement of net photosynthesis of three species of Plagiomnium mosses and its relation to the light and temperature , 1999 .

[63]  P. Reich,et al.  Convergence and correlations among leaf size and function in seed plants: a comparative test using independent contrasts. , 1999, American journal of botany.

[64]  William L. Wagner,et al.  Manual of the Flowering Plants of Hawai'i , 1999 .

[65]  P. Reich,et al.  Low-light carbon balance and shade tolerance in the seedlings of woody plants: Do winter deciduous and broad-leaved evergreen species differ? , 1999 .

[66]  F. Pugnaire,et al.  Handbook of Functional Plant Ecology , 1999 .

[67]  H. Rydin,et al.  Response of photosynthesis of Sphagnum species from contrasting microhabitats to tissue water content and repeated desiccation. , 1998, The New phytologist.

[68]  J. W. Bates Is 'life-form' a useful concept in bryophyte ecology? , 1998 .

[69]  M. Battaglia,et al.  Photosynthetic responses of , 1998 .

[70]  Peter R. Crane,et al.  The origin and early evolution of plants on land , 1997, Nature.

[71]  M. C. Davey Effects of short-term dehydration and rehydration on photosynthesis and respiration by Antarctic bryophytes , 1997 .

[72]  O. Lange,et al.  Water Relations and CO2 Exchange of Tropical Bryophytes in a Lower Montane Rain Forest in Panama , 1997 .

[73]  S. Caporn,et al.  The effects of long-term elevated ozone concentrations on the growth and photosynthesis of Sphagnum recurvum and Polytrichum commune. , 1996, The New phytologist.

[74]  Z. Tuba,et al.  Photosynthetic responses of a moss, Tortula ruralis, ssp. ruralis, and the lichens Cladonia convoluta and C. furcata to water deficit and short periods of desiccation, and their ecophysiological significance: a baseline study at present-day CO2 concentration. , 1996, The New phytologist.

[75]  S. K. Rice,et al.  The influence of water content and leaf anatomy on carbon isotope discrimination and photosynthesis in Sphagnum , 1996 .

[76]  I. Kovář Morphology and Anatomy , 1996 .

[77]  J. Grace,et al.  Deposition of fixed atmospheric nitrogen and foliar nitrogen content of bryophytes and Calluna vulgaris (L.) Hull. , 1995, Environmental pollution.

[78]  C. Delwiche,et al.  Phylogenetic Relationships of the "Green Algae" and "Bryophytes" , 1994 .

[79]  Bruce D. Clarkson,et al.  A manual of the flowering plants of Hawaii , 1992 .

[80]  E. Gorham Northern Peatlands: Role in the Carbon Cycle and Probable Responses to Climatic Warming. , 1991, Ecological applications : a publication of the Ecological Society of America.

[81]  N. Slack Bryophytes and ecological niche theory , 1990 .

[82]  D. Vitt Growth and production dynamics of boreal mosses over climatic, chemical and topographic gradients , 1990 .

[83]  H. Mooney,et al.  Stratification of tropical forests as seen in leaf structure , 1984, Tasks for vegetation science.

[84]  R. Longton,et al.  The biology of polar bryophytes and lichens: Contents , 1988 .

[85]  Thomas J. Givnish,et al.  Adaptation to Sun and Shade: a Whole-Plant Perspective , 1988 .

[86]  W. Oechel,et al.  COMPARATIVE PATTERNS OF NET PHOTOSYNTHESIS IN AN ASSEMBLAGE OF MOSSES WITH CONTRASTING MICRODISTRIBUTIONS , 1987 .

[87]  T. Simon The leaf-area index of three moss species (Tortula ruralis, Ceratodon purpureus, and Hypnum cupressiforme) , 1987 .

[88]  Thomas J. Givnish,et al.  On the economy of plant form and function. , 1988 .

[89]  Christopher B. Field,et al.  photosynthesis--nitrogen relationship in wild plants , 1986 .

[90]  W. Buck,et al.  Introduction to Bryology , 1986, The Canadian field-naturalist.

[91]  J. Duckett,et al.  The Experimental biology of bryophytes , 1985 .

[92]  J. Titus,et al.  Carbon Balance for Two Sphagnum Mosses: Water Balance Resolves a Physiological Paradox , 1984 .

[93]  W. Steere,et al.  New Manual of Bryology , 1984 .

[94]  M. Proctor Structure and ecological adaptation , 1984 .

[95]  M. Proctor Physiological Ecology: Water Relations, Light and Temperature Responses, Carbon Balance , 1982 .

[96]  T. Pócs Tropical Forest Bryophytes , 1982 .

[97]  R. S. Clymo,et al.  The Ecology of Sphagnum , 1982 .

[98]  K. Mägdefrau Life-forms of Bryophytes , 1982 .

[99]  P. Rundel,et al.  Water Relations and Photosynthetic Response of a Desert Moss , 1980 .

[100]  K. Mott,et al.  Carbon balance in relation to drying in four epiphytic mosses growing in different vertical ranges , 1979 .

[101]  M. Proctor,et al.  PHOTOSYNTHESIS, RESPIRATION AND WATER CONTENT IN BRYOPHYTES , 1979 .

[102]  P. Nobel,et al.  Introduction to biophysical plant physiology , 1974 .

[103]  E. Small Ecological Significance of Four Critical Elements in Plants of Raised Spagnum Peat Bogs , 1972 .

[104]  S. Vogel ''Sun leaves'' and ''shade leaves'' - Differences in convective heat dissipation. , 1968 .

[105]  Margaret C. Anderson Studies of the woodland light climate. II. Seasonal variation in the light climate , 1964 .

[106]  Margaret C. Anderson Studies of the Woodland Light Climate: I. The Photographic Computation of Light Conditions , 1964 .

[107]  D. I. Axelrod Adaptive Radiation. , 1962, Science.

[108]  Itsuo Miyata,et al.  Seasonal Variations of the Photosynthetic Efficiency and Chlorophyll Content of Epiphytic Mosses , 1961 .

[109]  K. Mellanby Physiological Ecology , 1958, Nature.

[110]  N. Parihar,et al.  An introduction to embryophyta , 1958 .

[111]  T. Hosokawa,et al.  THE DAILY COMPENSATION PERIOD AND VERTICAL RANGES OF EPIPHYTES IN A BEECH FOREST , 1957 .

[112]  E. Bartram Manual of Hawaiian Mosses , 1933 .

[113]  W. Watson XEROPHYTIC ADAPTATIONS OF BRYOPHYTES IN RELATION TO HABITAT. , 1914 .